Expression of α-farnesene and ethylene biosynthesis genes in relation to superficial scald in ‘White Winter Pearmain’ apple during storage

Ruirui Ding , Meiyan Wang

Horticulture Advances ›› 2026, Vol. 4 ›› Issue (1) : 13

PDF
Horticulture Advances ›› 2026, Vol. 4 ›› Issue (1) :13 DOI: 10.1007/s44281-026-00100-2
Research Article
research-article
Expression of α-farnesene and ethylene biosynthesis genes in relation to superficial scald in ‘White Winter Pearmain’ apple during storage
Author information +
History +
PDF

Abstract

Superficial scald is a physiological disorder in apples (Malus domestica). A two-year study was performed to investigate the relationship between α-farnesene, ethylene, and superficial scald in ‘White Winter Pearmain’ apples. In the first year, untreated apples were stored at 0°C for 12 weeks in an observational experiment. Results indicated that the expression of ethylene perception genes and α-farnesene metabolism-related genes correlated with the onset of superficial scald. The second year involved an interventional study, where apples were treated with 1.25 mM lovastatin (LOV), 1.4 mM ethephon, and 700 mM 1-methylcyclopropene (1-MCP), prior to storage at 0°C for 12 weeks. After the storage, both LOV and 1-MCP significantly reduced α-farnesene, conjugated trienols (CTols), and ethylene levels, as well as the incidence of superficial scald, whereas flesh firmness was enhanced. The combination of LOV and 1-MCP showed the most pronounced effect, as justified by pronounced alterations in the relative expression levels of α-farnesene- and ethylene-related genes (α-farnesene synthase, MdAFS; 1-aminocyclopropane-1-carboxylate synthase 1, MdACS1; MdACS4; MdACS5B; 1-aminocyclopropane-1-carboxylate oxidase 1, MdACO1; MdACO5; MdACO7; ethylene receptor 5, MdETR5; MdETRS2; and MdETR102) during storage. Correlation analysis showed a strong association between CTol content, ethylene production rate (EPR), and the expression of MdAFS, MdACS4, MdACS5A, MdACS5B, MdACS7, MdACO1, MdACO5, and MdETRS2 with superficial scald occurrence. These findings offer new insights into the molecular mechanisms by which α-farnesene and ethylene influence superficial scald development.

Keywords

Malus domestica / α-Farnesene and conjugated trienols / Ethylene production rate / Superficial scald / LOV and 1-MCP / Gene expression

Cite this article

Download citation ▾
Ruirui Ding, Meiyan Wang. Expression of α-farnesene and ethylene biosynthesis genes in relation to superficial scald in ‘White Winter Pearmain’ apple during storage. Horticulture Advances, 2026, 4(1): 13 DOI:10.1007/s44281-026-00100-2

登录浏览全文

4963

注册一个新账户 忘记密码

References

[1]

Alberts AWJ, Chen G, Kuron V, Hunt J, Huff C, Hoffman J, et al. . Mevinolin: a highly potent competitive inhibitor of hydroxymethylglutaryl-coenzyme A reductase and a cholesterol-lowering agent. Proc Natl Acad Sci U S A, 1980, 77: 3957-61

[2]

Alexander L, Grierson D. Ethylene biosynthesis and action in tomato: a model for climacteric fruit ripening. J Exp Bot, 2002, 53: 2039-55

[3]

Alwan TF, Watkins CB. Intermittent warming effects on superficial scald development of ‘cortland’, ‘delicious’ and ‘law rome’ apple fruit. Postharvest Biol Technol, 1999, 16: 203-12

[4]

Balaguera-Lopez HE, Espinal-Ruiz M, Rodriguez-Nieto JM, Herrera-Arevalo A, Zacarias L. 1-methylcyclopropene inhibits ethylene perception and biosynthesis: a theoretical and experimental study on cape gooseberry (Physalis peruviana L.) fruits. Postharvest Biol Technol, 2021, 174: 111476

[5]

Busatto N, Farneti B, Commisso M, Bianconi M, Iadarola B, Zago E, Ruperti B, Spinelli F, Zanella A, Velasco R, Ferrarini A, Chitarrini G, Vrhovsek U, Delledonne M, Guzzo F, Costa G, Costa F. Apple fruit superficial scald resistance mediated by ethylene inhibition is associated with diverse metabolic processes. Plant J, 2018, 93: 270-285

[6]

Bustin SA, Beaulieu JF, Huggett J, Jaggi R, Kibenge FS, Olsvik PA, Penning LC, Toegel S. MIQE precis: practical implementation of minimum standard guidelines for fluorescence-based quantitative real-time PCR experiments. BMC Mol Biol, 2010, 11 74

[7]

Bustin SA, Benes V, Garson JA, Hellemans J, Huggett J, Kubista M, Wittwer CT. The MIQE guidelines: minimum information for publication of quantitative real-time PCR experiments. Clin Chem, 2009, 55: 611-622

[8]

Chen PM, Varga DM, Mielke EA, Facteau TJ, Drake SR. Control of superficial scald on “d’anjou” pears by ethoxyquin: effect of ethoxyquin concentration, time and method of application, and a combined effect with controlled atmosphere storage. J Food Sci, 2010, 55: 167-70

[9]

Chye ML, Tan CT, Chua NH. Three genes encode 3-hydroxy-3-methylglutaryl-coenzyme a reductase in Hevea brasiliensis: hmg1 and hmg3 are differentially expressed. Plant Mol Biol, 1992, 19: 473-84

[10]

Clouse RM, Carraro N. A novel phylogeny and morphological reconstruction of the pin genes and first phylogeny of the acc-oxidases (acos). Front Plant Sci, 2014, 5: 296

[11]

DeEll JR, Murr DP, Porteous MD, Rupasinghe HPV. Influence of temperature and duration of 1-methylcyclopropene (1-MCP) treatment on apple quality. Postharvest Biol Technol, 2002, 24: 349-53

[12]

Ding RR, Che XK, Liu HZ, Du BY, Dong KT, Zhang YH. Effects of 1-MCP and storage temperature on transcription of mevalonate (MVA) enzyme genes of α-farnesene in ‘White Winter Pearmain’apples fruit. Sci Hortic, 2020, 259 108841

[13]

Ding R, Du B, Zhang Y. Conjugated trienols and programmed cell death are more closely related to superficial scald than reactive oxygen species in apple fruit stored at low temperature. Sci Hortic, 2019, 246: 597-603

[14]

Eisenreich W, Rohdich F, Bacher A. Deoxyxylulose phosphate pathway to terpenoids. Trends Plant Sci, 2001, 6: 78-84

[15]

Fan XT, Mattheis JP. Development of apple superficial scald, soft scald, core flush, and greasiness is reduced by MCP. J Agric Food Chem, 1999, 47: 3063-8

[16]

Gapper NE, Bai J, Whitaker BD. Inhibition of ethylene-induced α-farnesene synthase gene PcAFS1 expression in ‘d’anjou’ pears with 1-mcp reduces synthesis and oxidation of α-farnesene and delays development of superficial scald. Postharvest Biol Technol, 2006, 41: 225-33

[17]

Golding JB, Mcglasson WB, Wyllie SG. Relationship between production of ethylene and α-farnesene in apples, and how it is influenced by the timing of diphenylamine treatment. Postharvest Biol Technol, 2001, 21: 225-33

[18]

Gong Y, Song J, Palmer LC, Vinqvist-Tymchuk M, Fillmore S, Toivonen P, et al. . Tracking the development of the superficial scald disorder and effects of treatments with diphenylamine and 1-MCP using an untargeted metabolomic approach in apple fruit.. Food Chem (Oxf), 2021, 2: 100022

[19]

Ireland HS, Guillen F, Bowen J, Tacken EJ, Putterill J, Schaffer RJ, et al. . Mining the apple genome reveals a family of nine ethylene receptor genes. Postharvest Biol Technol, 2012, 72: 42-6

[20]

Isidoro N, Almeida DPF. α-farnesene, conjugated trienols, and superficial scald in “Rocha” pear as affected by 1-methylcyclopropene and diphenylarnine. Postharvest Biol Technol, 2006, 42: 49-56

[21]

Jemric T, Lurie S, Dumija L, Pavicic N, Hribar J. Heat treatment and harvest date interact in their effect on superficial scald of “granny smith” apple. Sci Hortic, 2006, 107: 155-63

[22]

Ju Z, Curry EA. Lovastatin inhibits α-farnesene synthesis without affecting ethylene production during fruit ripening in apples. J Am Soc Hortic Sci, 1999, 125: 105-10

[23]

Ju ZG, Curry EA. Lovastatin inhibits α-farnesene biosynthesis and scald development in “delicious” and “granny smith” apples and “d’anjou” pears. J Am Soc Hortic Sci, 2000, 125: 626-9

[24]

Jung SK, Watkins CB. Superficial scald control after delayed treatment of apple fruit with diphenylamine (DPA) and 1-methylcyclopropene (1-MCP). Postharvest Biol Tec, 2008, 50: 45-52

[25]

Karagiannis E, Michailidis M, Tanou G, Samiotaki M, Karamanoli K, Avramidou E, et al. . Ethylene –dependent and –independent superficial scald resistance mechanisms in ‘granny smith’ apple fruit. Sci Rep, 2018, 8: 11436

[26]

Larrigaudière C, Candan AP, Giné-Bordonaba J, Civello M, Calvo G. Unravelling the physiological basis of superficial scald in pears based on cultivar differences. Sci Hortic, 2016, 213: 340-345

[27]

Li T, Tan D, Yang X, Wang A. Exploring the apple genome reveals six acc synthase genes expressed during fruit ripening. Sci Hortic, 2013, 157: 119-23

[28]

Liao P, Hemmerlin A, Bach TJ, Chye ML. The potential of the mevalonate pathway for enhanced isoprenoid production. Biotechnol Adv, 2016, 34: 679-713

[29]

Lu X, Liu X, Li S, Wang X, Zhang L. Possible mechanisms of warming effects for amelioration of superficial scald development on ‘fuji’ apples. Postharvest Biol Tec, 2011, 62: 43-9

[30]

Lu X, Nock JF, Ma Y, Liu X, Watkins CB. Effects of repeated 1-methylcyclopropene (1-mcp) treatments on ripening and superficial scald of ‘cortland’ and ‘delicious’ apples. Postharvest Biol Tec, 2013, 78: 48-54

[31]

Lurie S, Lers A, Shacham Z, Sonego L, Burd S, Whitaker B. Expression of α-farnesene synthase afs1 and 3-hydroxy-3-methylglutaryl-coenzyme a reductase hmg2 and hmg3 in relation to α-farnesene and conjugated trienols in `granny smith’ apples heat or 1-mcp treated to prevent superficial scald. J Am Soc Hortic Sci, 2005, 130: 232-6

[32]

Pechous SW, Whitaker BD. Cloning and bacterial expression of a 3-hydroxy-3-methylglutaryl-coa reductase cdna (hmg1) from peel tissue of apple fruit. J Plant Physiol, 2002, 159: 907-16

[33]

Pechous SW, Whitaker BD. Cloning and functional expression of an (e,e)-α-farnesene synthase cdna from peel tissue of apple fruit. Planta, 2004, 219: 84-94

[34]

Ailed P, Armando M, Roxana UM, Javier B. Role of AtYap1 in the reactive oxygen species regulation of lovastatin production in Aspergillus terreus. Appl Microbiol Biotechnol, 2023, 107(4): 1439-1451

[35]

Moggial C, Hernández O, Pereiral M, Lobosl GA, Yuri JA. Effect of the cooling system and 1-mcp on the incidence of superficial scald in ‘granny smith’ apples. Chil J Agr Res, 2009, 69: 383-90

[36]

Rupasinghe HPV, Almquist KC, Paliyath G, Murr DP. Cloning of hmg1, and hmg2, cdnas encoding 3-hydroxy-3-methylglutaryl coenzyme a reductase and their expression and activity in relation to α-farnesene synthesis in apple. Plant Physiol Biochem, 2001, 39: 933-47

[37]

Rupasinghe HPV, Murr DP, Paliyath G, Skog L. Inhibitory effect of 1-mcp on ripening and superficial scald development in ‘mcintosh’ and ‘delicious’ apples. J Pomol Hort Sci, 2000, 75: 271-6

[38]

Sabbanamin R, Feygenberg O, Belausov E, Pesis E. Low oxygen and 1-mcp pretreatments delay superficial scald development by reducing reactive oxygen species (ros) accumulation in stored ‘granny smith’ apples. Postharvest Biol Technol, 2011, 62: 295-304

[39]

Schaffer RJ, Friel EN, Souleyre EJF, Bolitho K, Thodey K, Ledger S. A genomics approach reveals that aroma production in apple is controlled by ethylene predominantly at the final step in each biosynthetic pathway. Plant Physiol, 2007, 144: 1899-912

[40]

Tan LC, Zhao L, Liu XH, Deng CM, Guan ZZ. Antagonism of lovastatin on oxidative stress and apoptosis in primary rat hippocampal neurons induced by β-amyloid peptide. Chin J Pathol, 2017, 46(7): 6

[41]

Tong S, Zhao Y, Feng S. Effect of ethanol treatment in controlling superficial scald of “Red Star" Apples. Food Sci, 2002, 23: 124-127

[42]

Tsantili E, Gapper NE, Arquiza JMRA, Whitaker BD, Watkins CB. Ethylene and α-farnesene metabolism in green and red skin of three apple cultivars in response to 1-methylcyclopropene (1-mcp) treatment. J Agric Food Chem, 2007, 55: 5267-76

[43]

Varanasi V, Shin S, Johnson F, Mattheis JP, Zhu Y. Differential suppression of ethylene biosynthesis and receptor genes in ‘golden delicious’ apple by preharvest and postharvest 1-mcp treatments. J Plant Growth Regul, 2013, 32: 585-95

[44]

Watkins CB, Bramlage WJ, Cregoe BA. Superficial scald of `granny smith’ apples is expressed as a typical chilling injury. J Am Soc Hortic Sci, 1995, 120: 88-94

[45]

Whitaker B. Genetic and biochemical bases of superficial scald storage disorder in apple and pear fruits. Acta Hortic, 2013, 989: 47-60

[46]

Whitaker BD. Oxidation products of α-farnesene associated with superficial scald development in d’anjou pear fruit are conjugated trienols. J Agric Food Chem, 2007, 55: 3708-12

[47]

Xie X, Song J, Yan W, Sugar D. Ethylene synthesis, ripening capacity, and superficial scald inhibition in 1-mcp treated ‘d’anjou’ pears are affected by storage temperature. Postharvest Biol Technol, 2014, 97: 1-10

[48]

Xie XB, Zhao J, Wang Y. Initiation of ripening capacity in 1-mcp treated green and red ‘anjou’ pears and associated expression of genes related to ethylene biosynthesis and perception following cold storage and post-storage ethylene conditioning. Postharvest Biol Technol, 2016, 111: 140-9

[49]

Yang X, Song J, Campbell-Palmer L, Fillmore S, Zhang Z. Effect of ethylene and 1-mcp on expression of genes involved in ethylene biosynthesis and perception during ripening of apple fruit. Postharvest Biol Technol, 2013, 78: 55-66

[50]

Yang X, Song J, Du L, Forney C, Campbell-Palmer L, Fillmore S. Ethylene and 1-mcp regulate major volatile biosynthetic pathways in apple fruit. Food Chem, 2016, 194: 325-36

[51]

Yu J, Yan W. The combination of ethoxyquin, 1-methylcyclopropene and ethylene treatments controls superficial scald of ‘d’anjou’ pears with recovery of ripening capacity after long-term controlled atmosphere storage. Postharvest Biol Technol, 2017, 127: 53-9

[52]

Yuan K, Liu Q, Li B, Zhang L. Genomic structure and sequence polymorphism of e,e-alpha-farnesene synthase gene in apples ( malus domesticaborkh. ). Front Agr China, 2008, 2: 190-3

[53]

Zhou S, Cheng Y, Guan J. The molecular basis of superficial scald development related to ethylene perception and α-farnesene metabolism in ‘wujiuxiang’ pear. Sci Hortic, 2017, 216: 76-82

RIGHTS & PERMISSIONS

The Author(s)

PDF

0

Accesses

0

Citation

Detail

Sections
Recommended

/